An annotated catalog of the African primate genera Colobus and Procolobus (Cercopithecidae, Colobinae) in the collections of the American Museum of Natural History

26 Pages·2003·3 MB·English

by  American Museum of Natural History

Novitate MUSEUM
vitates
PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3399, 26 pp., 1 figure, 1 appendix March 27, 2003
An Annotated Catalog of the African Primate
Genera Colobus and Procolobus
(Cercopithecidae: Colobinae) in the Collections of
the American Museum of Natural History
RUTH O’LEARY!
ABSTRACT
The Department of Mammalogy of the American Museum of Natural History houses over
300 specimens of African colobus monkeys (genera Colobus and Procolobus). The collection
consists of red, olive, and black-and-white colobus (Procolobus badius, P. verus, Colobus
angolensis, C. guereza, C. polykomos, C. satanas, and C. vellerosus). Taxonomic notes, pelage
descriptions, and range are noted for each species and subspecies. For each specimen, the
following data are given: catalog number, date collected or recorded, sex, age, nature of spec-
imen, locality, name of collector, and measurements.
INTRODUCTION of the Democratic Republic of the Congo,
near Faradje. A map of the collecting sites
The American Museum of Natural History of the expedition was published by Allen
currently houses over 300 specimens of Af- (1925). The remainder of the colobus collec-
rican colobus monkeys. Herbert Lang and tion represents a spotty sampling from se-
James P. Chapin collected the majority of the lected geographic areas.
red colobus and over half of the black-and- The collection consists mostly of speci-
white colobus during the American Museum mens preserved as skins and skulls. There are
Congo Expedition (1909-1915). All of these also skeletons, casts, wet specimens, and
specimens were collected from the northeast mounted specimens present. The collection
' Scientific Assistant, Division of Vertebrate Zoology (Mammalogy), American Museum of Natural History. e-mail:
oleary @amnh.org.
Copyright © American Museum of Natural History 2003 ISSN 0003-0082
ws AMERICAN MUSEUM NOVITATES NO. 3399
contains the type series of Colobus langi (A1- reached and since this catalog is not meant
len, 1925), consisting of the holotype (Law- as a review of colobus taxonomy, I have fol-
rence, 1993) and two paratypes. This mate- lowed a conservative classification through-
rial is listed herein under the heading for out. Similarly, pelage descriptions refer only
Procolobus badius ellioti, currently regarded to specimens in the collection, not to the spe-
by Napier (1985) as a subjective senior syn- cies or subspecies as a whole. It is hoped that
onym. the information contained here will be a use-
The classification used in this catalog for ful resource for colobus studies and assist in
the black-and-white colobus follows Oates et the resolution of the taxonomy of these pri-
al. (1994) in separating these species at the mates.
generic level from the red and the olive co-
lobus, and in recognizing five species. At the
TAXONOMIC NOTES
subspecific level, the classification used by
Napier (1985) is followed, except where not- In 1781, Pennant published the first de-
ed. All five species of black-and-white co- scription of two colobus species—the ‘“‘full-
lobus are represented in the museum collec- bottom monkey” (the black-and-white colo-
tion. These are Colobus angolensis, C. guer- bus) and the “‘bay monkey” (the red colo-
eza, C. polykomos, C. satanas, and C. vel- bus). The descriptions were based on one
lerosus. specimen of each species in the Leverian
The taxonomy of the red colobus is unsta- Museum, London, brought from Sierra Le-
ble and has yet to be fully resolved. The clas- one. The type specimens were subsequently
sification used in this catalog is conservative lost during the dispersal of the Leverian col-
and follows Oates et al. (1994) at the generic lection. Later authors (e.g., Rochebrune,
level. The red colobus is recognized as a sub- 1886-1887) suggested that Pennant’s de-
genus, Piliocolobus of the genus Procolobus. scription and figures were unreliable. These
At the specific level, the catalog follows Na- factors, among others, contributed to the en-
pier (1985) in recognizing only two species, suing confusion regarding colobus nomen-
P. badius and P. kirkii. Procolobus kirkii is clature and classification.
considered by many authors to be a valid The taxonomy of the African colobus has
species, and although P. badius may in fact undergone a number of changes in the more
be comprised of several species, it is recog- than two hundred years since Pennant’s pub-
nized here as a single species. This arrange- lication. The debates center on the number of
ment conflicts with Groves (1993), who rec- genera that should be used to represent the
ognizes four species, and others (Dandelot, African colobus and the number of species
1971; Delson et al., 1982). that should be used to reflect the diversity of
Seven recognized subspecies or popula- the red colobus and the black-and-white co-
tions of red colobus are represented in the lobus.
collection: Procolobus (Piliocolobus) badius The black-and-white, red, and olive colo-
badius, P. b. bouvieri, P. b. ellioti, P. b. ous- bus are separated at the generic and subge-
taleti, P. b. rufomitratus, P. b. tephrosceles, neric levels. Initially, all three were consid-
and P. b. tholloni. A second species of red ered to be members of a single genus (Mar-
colobus, P. kirkii, is not represented in the tin, 1841) but subsequent research has re-
AMNH collection. vealed significant differences between each
Only one species of olive colobus is rec- group. Pocock (1936) examined the external
ognized, Procolobus (Procolobus) verus, of characters of the red colobus and compared
which there are two specimens in the collec- the results to work he had carried out pre-
tion. viously on black-and-white colobus (Pocock,
The primary purpose of this catalog is to 1926). His data led him to suggest that the
record and present the data on all the colobus two should be distinguished generically. Hill
specimens in the AMNH. Recent reviews of (1952) conducted studies of the external and
colobus taxonomy have raised several sub- visceral anatomy of the olive colobus and
species to specific status (e.g., Groves, 2001). concluded that his findings suggested a rea-
However, since no consensus has_ been son to separate verus (the olive colobus)
2003 O’LEARY: CATALOG OF COLOBUS AND PROCOLOBUS 3
from badius (the red colobus). Many simi- agreement yet on which of these should be
larities between the red and olive colobus assigned specific or subspecific status. It is
were being uncovered at this time and there generally agreed that the Zanzibar red colo-
was a need to represent these shared features bus, P. kirkii, is sufficiently distinct morpho-
in the taxonomy. Hill and Booth (1957) sug- logically and ecologically from the other red
gested uniting Procolobus (the olive colo- colobus to be distinguished at the specific
bus) and Piliocolobus (the red colobus) with- level (Verheyen, 1962; Napier, 1985).
in the genus Procolobus based on their dis- A comparison of classifications for the red
tinctive shared features. These features in- colobus can be found in Oates et al. (1994),
clude sexual swelling in the female, the while Groves (2001) contains a more recent
presence of a perineal organ in the male, sep- classification. The number of species used to
arate ischial callosities in the female, and describe the group varies considerably. Rahm
small larynx (Pocock, 1936; Hill, 1952; Hill (1970) recognized only one species, Dande-
and Booth, 1957; Kuhn, 1967, 1972). All of lot (1971) eight species, and Napier (1985)
these features were observed to be common two species. More recently, Colyn (1991)
to the red and the olive colobus, and to dis- suggested that the subspecies ellioti may ac-
tinguish both from the black-and-white co- tually consist of four distinct subspecies (lan-
lobus. In addition, Clutton-Brock (1974) and gi, lulindicus, foai, and semlikiensis) together
Struhsaker and Oates (1975) noted signifi- with subspecific hybrids. Groves (2001) rec-
cant differences in the behavior and ecology ognized nine species.
of the red and the black-and-white colobus There has been little resolution to this sub-
in the Kibale Forest, Uganda, which also ject, however, and Oates et al. (1994) ob-
suggested that the two groups might not be served that an appropriate classification may
members of the same genus. not be reached until further comprehensive
Currently, Oates et al. (1994) and others research is carried out.
(Brandon-Jones, 1984; Strasser and Delson, There is only one recognized species of
1987) agree that the red and the olive colo- olive colobus, Procolobus (Procolobus) ve-
bus monkeys are more closely related to each rus.
other than either is to the black-and-white co-
lobus. These authors recognize two genera—
FORMAT OF THE CATALOG
Colobus, for the black-and-white colobus
and Procolobus to represent the red and the The African colobus collection in the
olive colobus. Procolobus is further subdi- AMNH is arranged according to genus and
vided into two subgenera, Piliocolobus (red) subgenus; the two genera are Colobus, the
and Procolobus (olive). This follows an ar- black-and-white colobus, and Procolobus,
rangement suggested by Hill and Booth including the subgenera Procolobus (the ol-
(1957). It is also the arrangement followed ive colobus) and Piliocolobus (the red colo-
in this catalog. bus). This catalog followed the format used
The taxonomy of the black-and-white co- in An annotated catalogue of Malagasy pri-
lobus has been worked out with some reso- mates... (Buettner-Janusch and Tattersall,
lution. At least four and, more recently, five 1985). For each level of classification, infor-
distinct species are recognized (Oates et al., mation was given under the following head-
1994; Groves, 2001). The elevation of vel- ings:
lerosus to specific status was suggested by TAXONOMIC NOTES: As discussed above,
Dandelot (1971) and investigated further by due to the unstable nature of red colobus sys-
Oates and Trocco (1983), whose findings led tematics in particular, there are a number of
them to conclude that vellerosus is a distinct different classifications proposed at both the
species. specific and subspecific level. Under this
The taxonomy of the red colobus at spe- heading, alternative classifications of the
cific level has yet to be resolved with satis- groups and the opinions of different authors
faction. The same 14 forms are recognized are presented.
by many authors (Rahm, 1970; Dandelot, PELAGE Notes: Different formats of de-
1971; Napier, 1985). However, there is no scription were used for the red and the black-
AMERICAN MUSEUM NOVITATES NO. 3399
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2003 O’LEARY: CATALOG OF COLOBUS AND PROCOLOBUS 5
and-white colobus in order to best describe tag noted a specimen as an embryo. Follow-
features of the pelage of each group. For the ing Napier (1985), an adult was defined as
red colobus, a description of coat coloration having complete permanent dentition. Adults
was given for each subspecies if relevant, included individuals whose canines were not
followed by the distinguishing characteristics fully grown. An infant was defined as having
of the pelage of each subspecies. Wherever entirely deciduous dentition, without M1
possible, the pelage description followed this erupted. A juvenile included all stages of de-
order: head, cheeks, nape, shoulders, back, velopment between adult and infant, as well
sides, underparts, forelimbs, hind limbs, tail. as individuals with nonerupted M3. For some
In the case of the black-and-white colobus, of the specimens without skulls, the age was
general pelage descriptions were outlined for inferred from the skin size. In these cases,
each species. The key formatted by Napier the age was enclosed in parentheses.
(1985) was used to distinguish subspecies in Description: Skull, skin, and skeleton were
C. angolensis and C. guereza. Napier’s key noted when present. Skull included cranium
used the most appropriate features of the pel- and mandible; specimens with anything less
age to describe the variation in each subspe- were noted as partial skull. Skin denoted
cies. AS mentioned previously, the pelage de- whole skin, either flat or mounted.
scriptions referred only to the specimens in Locality: Data were taken from field tags,
the AMNH collection. and the localities were verified in a modern
RANGE OF SPECIMENS: The distribution of
atlas. Field notes and gazetteers of the orig-
the AMNH specimens was noted for each inal collectors were also used as references
group, followed by the range of the subspe- for locality information. When available, the
cies as a whole according to the literature. altitude of the locality was also included. Al-
SPECIMENS: A species list was given in the
len (1925) included a map of localities from
appendix for each subspecies. the Lang and Chapin expedition which was
useful for many of the specimens in this col-
SPECIES LISTS lection. If information was not available, or
Eight columns were used to describe each if the specimen was taken from a zoo, “‘No
taxon in this collection, under the following Data”’ was entered in this column.
headings: In the case of one species and three sub-
Catalog Number: The number assigned by species (i.e., Colobus polykomos, C. ango-
the museum to each specimen. lensis cottoni, C. guereza guereza, and C. g.
Date: This was the date of collection of kikuyuensis) specimens without exact locali-
wild-caught specimens or, in the case of zoo ty data were included in the list. This was
specimens, the date on which they were re- done when the pelage characters correspond-
corded in the permanent catalog of the ed accurately with the descriptions given by
AMNH. Napier (1985) and Dandelot (1971) and with
Sex and Age: Information on sex was tak- the other specimens of that species or sub-
en from the field tag and confirmed from the species in the collection.
actual specimen whenever possible. Three The Democratic Republic of the Congo
age classes were recorded: adult, juvenile, was abbreviated throughout as DRC. This
and infant. In four instances, the collector’s country was known as the Belgian Congo at
<
Fig. 1. Collection localities in the Congo Basin. Hf represent localities where black-and-white co-
lobus were collected, L] represent localities where red colobus were collected. Localities where both
red, and black-and-white colobus were collected are marked with a Ll. The localities shown on the map
are as follows: Congo (Brazzaville): 1. Makoua, 2. Ouesso. Democratic Republic of the Congo: 3.
Abawe, 4. Akenge, 5. Angumu, 6. Avakubi, 7. Bafuka, 8. Bafwabaka, 9. Beni, 10. Bolobo, 11. Faradje,
12. Gamangui, 13. Kananga, 14. Lukolela, 15. Medje, 16. Niapu, 17. Poko, 18. Risimu, 19. Rutshuru,
20. Ukaturaka, 21. Vankerckhovenville, 22. Yakuluku. Uganda: 23. Kibale Forest, Kanyawara, 24.
Kibale Forest, near Dubona camp, 25. Malabigambo Forest.
6 AMERICAN MUSEUM NOVITATES NO. 3399
the time most of the specimens were col- tinctive pelage characters; for example, C.
lected. It was later called Zaire. The Republic angolensis has shoulder hairs or epaulettes,
of Congo was noted as Congo (Brazzaville) which readily distinguish it from other Co-
to distinguish the two countries. This country lobus species. The various forms of these
was formerly known as French Congo. features were also used to identify subspe-
Remarks: In this column the name of the C18:
collector was listed, along with the original RANGE OF SPECIMENS: The specimens of
field number of the specimen. All zoo spec- the genus Colobus in the museum were col-
imens were from the NYZS (New York Zoo- lected from Liberia, Ivory Coast, Cameroon,
logical Society) and were noted accordingly. and Gabon in the west and across the conti-
Measurements: The measurement column nent from Congo (Brazzaville) to Ethiopia
lists data taken from the collector’s field tag and south to Tanzania. The entire range of
and includes total length, tail, hind foot, and black-and-white colobus is more widespread,
ear length. The measurements were given in from Guinea Bissau in the west, including
millimeters for most specimens and were in- northeastern Angola and possibly Zambia to
cluded in parentheses in inches if originally the south and Sudan and the Central African
measured as such by the collector. Republic to the north (Dandelot, 1971; Oates
Specimens taken from zoos or without suf- and Trocco, 1983; Napier, 1985).
ficient data to enable identification at the sub-
specific level are included at the end of the Colobus angolensis Sclater, 1860
appendix, nos. 22-25.
TAXONOMIC NOTES: Rahm (1970) and Na-
GAZETTEER pier (1985) recognized eight subspecies.
Dandelot (1971) recognized only six, syn-
Geographic coordinates for the majority of
onymizing adolfifriederici with ruwenzorii
the collecting localities were listed in the
and palliatus with sharpei. Groves (2001)
gazetteer. A map of the central African lo-
recognized the same six subspecies as Dan-
calities (fig. 1) included the distributions of
delot (1971) and included an unnamed sub-
the specimens collected from that area.
species from the northern part of the Mahale
mountains, east of Lake Tanganyika. This
GENUS COLOBUS ILLIGER, 1811
catalog followed Napier (1985) in maintain-
TAXONOMIC NOTES: The genus Colobus ing the separate status of adolfifriederici and
represents only the black-and-white colobus ruwenzorii and of palliatus and sharpei.
monkeys. For some time, four species were PELAGE NorTEs: The long white shoulder
recognized: Colobus angolensis, C. guereza, hairs, which are continuous with the white
C. polykomos, and C. satanas. However, hairs of the cheeks and throat, distinguish
Oates and Trocco (1983) suggested elevating Colobus angolensis from other Colobus spe-
C. polykomos vellerosus to specific status cies. Some subspecies have a white brow
based on their study of vocalizations, cranial band. There is no white mantle and the tail
dimensions, coat pattern, and geographical is untufted. The proximal portion of the tail
distribution. Following their findings, five is black and the distal portion is white or
species of black-and-white colobus are rec- grizzled. The proportion of the grizzled part
ognized in this catalog. of the tail varies in length according to sub-
PELAGE NOTEs: The five species of the ge- species. Napier (1985) used features such as
nus Colobus were easily identified by their the brow band, epaulettes, pubic patch, and
black-and-white or all black (C. satanas) tail coloration to identify subspecies; that
coats. The black-and-white colobus are large format is followed here.
monkeys and have long tails which in C. RANGE OF SPECIMENS: The Colobus ango-
guereza are tufted. The infant pelage con- lensis specimens in the AMNH were col-
trasts markedly with the adult pelage, being lected from areas across northern DRC,
completely white at birth in all species ex- Uganda, and southwestern Tanzania. Colo-
cept C. satanas, whose coat is brown at birth bus angolensis is distributed across mainland
(Oates et al., 1994). Each species has dis- Africa from southern Congo (Brazzaville),
2003 O’LEARY: CATALOG OF COLOBUS AND PROCOLOBUS zs
northeastern Angola, DRC (excluding the and Lomani Rivers to the southern border of
northwest and southeast regions), Rwanda, Katanga (Dandelot, 1971).
Uganda, and Tanzania and is also found at SPECIMENS: Appendix 1, no. 2.
the Kenyan coast (Dandelot, 1971; Napier,
1985; Groves, 2001). Colobus angolensis cottoni Lydekker, 1905
PELAGE NOTES: Brow band absent, epau-
Colobus angolensis adolfifriederici
lettes short, not very profuse, pubic region
Matschie, 1914b
entirely black. Tail black at base, grizzled for
the remaining 80%.
TAXONOMIC NOTES: Dandelot (1971) con-
RANGE OF SPECIMENS: All of the specimens
sidered adolfifriederici to be synonymous
for which there are locality data available are
with ruwenzorii. He proposed that the only
from northeastern DRC, which is the range
distinction between the two forms is in pel-
age density; adolfifriederici, living at higher of the subspecies noted by Rahm (1970) and
Dandelot (1971). The last two specimens in
altitudes than ruwenzorii, has a longer pel-
the list do not have locality data. However,
age. Hull (1979) also united adolfifriederici
Allen (1925) included them as C. a. cottoni,
with ruwenzorii based on craniometric stud-
and the pelage markings of the two speci-
ies. Rahm (1970), however, did not agree,
contesting that there are significant differenc- mens are comparable to the rest of the col-
lection.
es between the two in shoulder hairs and in
SPECIMENS: Appendix 1, no. 3.
tail coloration. Napier (1985) also noted dis-
tinctions in the pelage between the two
forms, including less profuse epaulettes in Colobus angolensis ruwenzorii
adolfifriederici (mentioned by Rahm, 1970) Thomas, 1901
and a broader white pubic patch in ruwen- TAXONOMIC NOTES: See notes for C. a.
zorii. Groves (2001) united the two subspe-
adolfifriederici.
cies. Given the uncertainty, adolfifriederici
PELAGE NOTES: There are two skin speci-
and ruwenzorii are treated in the catalog as
mens of ruwenzorii in the museum collec-
separate subspecies in line with following a tion, only one of which is complete. The fol-
conservative classification.
lowing description is based on that skin.
PELAGE NOTES: There is only one skin
Long, dense pelage. Narrow brow band, long
specimen of adolfifriederici in the collection.
epaulettes, broad grizzled pubic patch. Tail
Narrow brow band, epaulettes less profuse
almost all black, distal 5% grizzled.
than those of C. a. angolensis. Pubic patch
RANGE OF SPECIMENS: Ruwenzori Moun-
small and grizzled. Tail almost entirely black,
tains. This is the range of the subspecies ac-
distal 15% grizzled.
cording to Rahm (1970) and Napier (1985).
RANGE OF SPECIMENS: Southern Uganda.
Dandelot (1971) and Groves (2001) extended
The distribution of the subspecies is eastern
the range of ruwenzorii, as they have united
DRC, Rwanda, and southwestern Uganda it with adolfifriederici.
(Napier, 1985). Rahm (1970) extended the
SPECIMENS: Appendix 1, no. 4.
distribution to Minziro forest in Tanzania,
which is just over the border with Uganda.
Colobus angolensis sharpei Thomas, 1902
SPECIMENS: Appendix 1, no. 1.
TAXONOMIC NOTES: Dandelot (1971) unit-
Colobus angolensis angolensis Sclater, 1860 ed sharpei with palliatus for the same reason
he united adolfifriederici with ruwenzorii; he
PELAGE NOTES: Brow band absent, epau- argued that increased pelage density in shar-
lettes long. Approximately 25% of the distal pei is the only distinction between the two.
portion of the tail is grizzled. Groves (2001) also untied the two. Napier
RANGE OF SPECIMENS: Western and south- (1985), however, noted various other differ-
ern DRC. The subspecies is found in north- ences in the pelage of the two forms, and
eastern Angola and DRC (Rahm, 1970; Dan- Hull (1979) suggested keeping sharpei and
delot, 1971) from the left bank of the Congo palliatus separate pending further study. The
8 AMERICAN MUSEUM NOVITATES NO. 3399
two are considered separate subspecies in RANGE OF SPECIMENS: Eastern Ethiopia.
this catalog. Ethiopia, northeast of Rift Valley, Galla
PELAGE NOTES: Long pelage, distinct, country (Dandelot, 1971).
broad white brow band. Long epaulettes, Note: The colobus monkeys collected by
white, broad pubic patch. Distal 25% of tail T. D. Carter at Mount Ansha include what
grizzled. appear to be specimens of both C. g. gallar-
RANGE OF SPECIMENS: Southern Tanzania. um and C. g. guereza. Napier (1985) and
Rahm (1970) noted the distribution as north Dandelot (1971) distinguished the two sub-
of Lake Malawi, and Rungwe, Poroto, Liv- species by the color of their tail base. Al-
ingstone, and Uzungwa Mountains, Tanza- though the tail base of the specimens of C.
nia. g. gallarum is somewhat grizzled under-
SPECIMENS: Appendix 1, no. 5.
neath, it is distinguishable from the speci-
mens of C. g. guereza. Yalden et al. (1977)
Colobus guereza Riippell, 1835 expressed dissatisfaction with the basis of the
TAXONOMIC NOTES: Nine subspecies were separation, stating that the variation in tail
recognized by Schwarz (1929) and Rahm color may be clinal in nature. Furthermore,
it is not clear if the distribution or range of
(1970). Dandelot (1971) recognized only six,
synonymizing percivali with guereza and the two subspecies overlaps. Rahm (1970)
uellensis and dodingae with occidentalis. included a map of the distribution of all the
Hull’s study (1979) of craniometric charac- subspecies of Colobus. In this map, the dis-
ters led him to recognize eight subspecies, tribution of C. g. gallarum does not extend
uniting uellensis with occidentalis. Groves as far south or west as Dandelot (1971) sug-
(2001) and Napier (1985) concurred with this gested. Yalden et al. (1977) stated that there
arrangement, and that classification is fol- appears to be no geographic boundaries be-
lowed here. tween the two forms to separate them or to
PELAGE NOTEs: The distinguishing features prevent dispersal.
of the pelage of C. guereza are the long Furthermore, the location of Mt. Ansha
white mantle and the long flowing tail, which could not be determined exactly. It is north of
ends in a white tuft. This species is also not- Mt. Cacca (Kaka), and the province Carter
ed for its wide white ring around the callos- (collector) called Arussi is now called Arsi
ities. The features used by Dandelot (1971) (Yalden, personal commun.) While it is ac-
and Napier (1985) to distinguish between knowledged that the same locality is noted for
subspecies are the length of the mantle, color two subspecies, they were separated in the
of tail base, and description of the white tail collection according to pelage differences.
tuft. SPECIMENS: Appendix 1, no. 6.
RANGE OF SPECIMENS: The Colobus guer-
eza specimens in the AMNH were collected
Colobus guereza guereza Ritippell, 1835
from localities in Congo (Brazzaville), north-
eastern DRC, Uganda, Kenya, and Ethiopia. TAXONOMIC NOTES: Dandelot (1971) unit-
Napier (1985) and Dandelot (1971) recorded
ed percivali with guereza, while other au-
the range of the species from eastern Nigeria
thors (Rahm, 1970; Hull, 1979; Napier, 1985;
through Cameroon, northern Gabon, Congo
Groves, 2001) maintained the two as separate
(Brazzaville), eastwards through northern
subspecies.
DRC, Central African Republic, southern
PELAGE Notes: Long mantle, covering
Sudan and Ethiopia, and southwards to Ken-
base of tail, which is grizzled. White tuft
ya, Uganda, and northern Tanzania.
comprises half of the tail.
RANGE OF SPECIMENS: Western Ethiopia.
Colobus guereza gallarum Neumann, 1902
Groves (2001) reported the range from for-
PELAGE NoTEs: Long mantle, covering tail ested areas of the Ethiopian highlands, ex-
base which is black, grizzled underneath. tending into lowland forests along the Omo
White tuft is almost half of the total tail River and in the Blue Nile gorge.
length. SPECIMENS: Appendix 1, no. 7.
2003 O’LEARY: CATALOG OF COLOBUS AND PROCOLOBUS 9
Colobus guereza kikuyuensis pier, 1985) recognized three subspecies (po-
Loénnberg, 1912 lykomos, dollmani, and vellerosus). Dandelot
(1971), however, noted that vellerosus might
PELAGE NOTES: Very long mantle, cover-
actually be a distinct species. More recently,
ing up to a quarter of the tail. Base of tail
Oates and Trocco (1983) suggested raising C.
black, white tuft voluminous, forming about
polykomos vellerosus to specific status based
two-thirds of the length of the tail.
on vocalization and other analyses. Oates et
RANGE OF SPECIMENS: Southwestern Ken-
al. (1994) followed Oates and Trocco (1983)
ya. The subspecies occurs at Ngong Escarp-
in recognizing vellerosus as a separate spe-
ment, Mount Kenya, and the Aberdare Range
cies. Booth (1958) and Rahm (1970) sug-
(Dandelot, 1971; Groves, 2001).
gested that the intermediate form, dollmani,
SPECIMENS: Appendix 1, no. 8.
might in fact be a hybrid swarm, a theory
Colobus guereza matschiei investigated by Groves et al. (1993). They
Neumann, 1899 conclude that dollmani is in fact a hybrid and
that polykomos and vellerosus should be rec-
PELAGE Notes: Long mantle, covering
ognized as distinct species. In this catalog, I
base of tail. Tail base black, tail tuft full,
follow Groves et al. (1993) and Groves
comprising about one-third of total length.
(2001) in recognizing vellerosus and poly-
RANGE OF SPECIMENS: Eastern Uganda and
komos as separate species. There are no spec-
western Kenya. Dandelot (1971) noted the
imens labeled C. p. dollmani in the AMNH
range of the subspecies as Kenya, west of
collection.
Rift Valley, Mau Escarpment, and Mount El-
PELAGE NOTES: Gray brow band, gray long
gon.
epaulettes. Tail white from base to tip and
SPECIMENS: Appendix 1, no. 9.
untufted. Callosities bordered by narrow ring
of white hair.
Colobus guereza occidentalis
RANGE OF SPECIMENS: Liberia and south-
(Rochebrune, 1886—1887)
ern Ivory Coast. The range of the species is
TAXONOMIC NOTES: Rahm (1970) main- from Guinea to Sassandra River, Ivory
tained occidentalis and uellensis as separate
Coast (Napier, 1985; Groves, 2001). Rahm
subspecies, while Dandelot (1971), Napier (1970) included Gambia in the range. How-
(1985), and Groves (2001) treated uellensis ever, Oates and Trocco (1983) suggested
as a synonym of occidentalis. Hull’s cranio-
that it is unlikely that C. polykomos is pre-
metric studies (1979) confirmed the view that sent in Gambia. Schwarz (1929) stated that
the two are synonymous. the type of C. leucomeros was almost cer-
PELAGE NOTES: Short mantle, not extend- tainly not from “‘River Gambia’’. Oates and
ing beyond base of tail, which is black. There
Trooco (1983) were unaware of other spec-
is variation among the specimens in the imens collected in Gambia and therefore ex-
length of the tail tuft—from one-fifth to one-
cluded Gambia from the range of C. poly-
third of the total length. komos.
RANGE OF SPECIMENS: There are three main SPECIMENS: Appendix 1, no. 11.
collection localities for the specimens of oc-
cidentalis here: southern and western Ugan-
Colobus satanas Waterhouse, 1838
da, northeastern DRC, and western Congo
(Brazzaville). The subspecies is found across TAXONOMIC NOTES: Dandelot (1971) and E1-
Africa from eastern Nigeria, southern Cam- sentraut (1973) agreed that the subspecies met-
eroon, Gabon, and northern DRC to Uganda ternichi was in fact based on partial albinos.
and southwestern Sudan (Napier, 1985; Eisentraut (1973) also suggested that the main-
Groves, 2001). land population may be subspecifically distinct
SPECIMENS: Appendix 1, no. 10. from the island population on Bioko.
PELAGE NOTES: Coat entirely black.
Colobus polykomos (Zimmermann, 1780)
RANGE OF SPECIMENS: The specimens in
TAXONOMIC NOTES: Until recently, most this collection were taken from southwestern
authors (Rahm, 1970; Dandelot, 1971; Na- Cameroon and northwestern Gabon. The
10 AMERICAN MUSEUM NOVITATES NO. 3399
subspecies occurs in Bioko, southwestern range extends from Liberia and Ivory Coast
Cameroon, Equatorial Guinea, Gabon, and in the west, across DRC and Uganda, with a
northwestern Congo (Brazzaville) (Rahm, few specimens from Tana River, Kenya.
1970; Carpaneto 1995). Dandelot (1971) stated that the red colobus
SPECIMENS: Appendix 1, no. 12. monkeys are to be found across the African
continent from Senegal in the west to the is-
Colobus vellerosus (1. Geoffroy, 1834) land of Zanzibar in the east.
TAXONOMIC NOTEs: As already noted, this
Species was previously considered a subspe- Procolobus (Piliocolobus) badius
(Kerr, 1792)
cies of C. polykomos, due, according to
Dandelot (1971), “‘to a craze for simplifi-
TAXONOMIC NOTES: This species may ac-
cation’”’.
tually be comprised of as many as eight spe-
PELAGE NOTES: The six skins in the
cies (Dandelot, 1971).
AMNH collection are not complete and con-
PELAGE NoTEs: All of the P. badius sub-
sist only of the trunk of the body up to the
species in this collection have a dark brown
forearms and down to the thighs. The griz-
or black band across the brow extending to
Zled patches on the thighs, the distinguishing
the ears. They are similar in body size and
feature of this species, are visible on the
can be easily distinguished by their respec-
skins.
tive pelage colors which have variable
RANGE OF SPECIMENS: There were no lo-
amounts of black, brown, and red.
cality data for the specimens of vellerosus in
RANGE OF SPECIMENS: The seven subspe-
this collection. The range of the subspecies
cies of P. badius in this collection are rep-
is from Bandama River, Ivory Coast, to west-
resentative of populations in north-central Li-
ern Nigeria (Dandelot, 1971; Rahm, 1970;
beria, western Ivory Coast, western DRC,
Groves, 2001).
northeastern DRC (the majority of the spec-
SPECIMENS: Appendix 1, no. 13.
imens was from this region), Uganda, and
Tana River, Kenya. As noted for the subge-
GENUS PROCOLOBUS, SUBGENUS
nus Piliocolobus, the actual range of these
PILIOCOLOBUS ROCHEBRUNE,
monkeys extends across the African conti-
1886-1887
nent.
TAXONOMIC NOTES: The subgenus Pilio-
colobus of the genus Procolobus represents Procolobus (Piliocolobus) badius badius
the red colobus monkey. (Kerr, 1792)
PELAGE NOTEs: The red colobus have been
described as large, slender monkeys with PELAGE NOTES: Two rich colors are seen
long, untufted tails (Napier, 1985). There is in this subspecies—black and maroon. Nei-
no sexual dimorphism in the coat coloration. ther color lessens in intensity throughout the
The pelages of infants and adults differ coat, as is often seen in the varying shades
somewhat in hue but not as markedly as in of brown observed in the pelages of other
the black-and-white colobus. The amount of subspecies. Black head, maroon cheeks.
red coloration in the pelages of these mon- Nape black with maroon shoulders. Back and
keys varies considerably among subspecies. rump black. Sides maroon. Upper arms
Some of the subspecies (e.g., P. b. tholloni) black, lower arms and hands maroon. Thighs
are strikingly red in color. Others exhibit lit- black, legs and feet maroon. Tail black from
tle red color; rather, their pelts are drab base to tip.
brown, as seen in P. b. oustaleti. Napier RANGE OF SPECIMENS: Liberia to Ivory
(1985) has published useful descriptions and Coast. The range of the subspecies is from
pelage notes on this group. Sierra Leone, eastern Guinea, to western Ivo-
RANGE OF SPECIMENS: The range of the red ry Coast. Eastern limit of their range is
colobus monkeys in the AMNH collection is thought to be the Bandama River (Dandelot,
somewhat limited, as only 7 of the approxi- 1971; Groves, 2001).
mate 14 subspecies are represented. The SPECIMENS: Appendix 1, no. 14.